vertebrate fauna n.l. mckenzie, lk.rolfe and w.k. …museum.wa.gov.au/sites/default/files/4....

35
VERTEBRATE FAUNA N.L. McKenzie, lK. Rolfe and W.K. Youngson Background The philosophies underlying our approach to this vertebrate survey, the sampling strategy adopted, and the actual methods we employed are described in Biological Surveys Committee of Western Australia (1984). Specimens representing the array of reptile and mammal species encountered in the Sandstone-Sir Samuel and Laverton-Leonora Study Areas during our survey have been lodged in the Western Australian Museum within the following series of register numbers: R62735- 62872, 70827-70877, 78545-78585, 79000, R114956-1149570, 114983-114985; M20395, 20396, 20404-20409, 20420, 20421, 20424, 20425, 20427, 20428, 20446-20448, 20461- 20463, 20815, 20816, 20821-20823, 20825-20830, 20832-20834, 20837, 20838, 20840, Table 2 Faunal survey of the Sandstone-Sir Samuel (Wanjarri) and Laverton-Leonora (Erlistoun) Study Areas: sampling dates and methods*. 1. Wanjarri Survey Area. Fauna 14-21 Feb, 14-20 May, 24-29 Aug, Lithological Code** 1979 1980 1981 Surface 1W-01 Q(M,B,R) Q(M,B,R) Q(M,B,R) Qpv/Qqz 1W-02 Q(M,B,R) Q (M,B,R) Q(M,B,R) Qqc 1W-03 Q(M,B,R) Q (M,B,R) Q(M,B,R) Qps/Qqz 1W-04 Q(M,B,R) Q(M,B,R) Q(M,B,R) Czo+Qpv lW-OS Q(M,B,R) Q (M,B,R) Q(M,B,R) Qqz 1W-06 F,B,R F,M,B,R F,M,B,R Qps 1W-07 F,B,R F,B,R Qps 1W-08 F,B,R F,B,R Qps 2. Erlistoun Survey Area. Fauna 22-28 Feb, 8-13 May, 18-23 Aug, Lithological Code** 1979 1980 1981 Surface lE-01 Q(M,B,R) Q(M,B,R) Q(M,B,R) Qpk 1E-02 Q(M,B,R) Q(M,B,R) Q(M,B,R) Qrm lE-03 Q(M,B,R) Q(M,B,R) Q(M,B,R) Qrd 1E-04 Q(M,B,R) Q(M,B,R) Q(M,B,R) Qps/Qqz lE-OS Q(M,B,R) Q(M,B,R) Q(M,B,R) Qpv lE-06 F,B,R F,B,R Qps * M = Mammal trapline (9 Elliotts, 9 Break-backs, 3 Cages) B = Bird searches R = Reptile searches F = Fenced pit line of six pits (4 shallow, 2 deep) with ca. 50 m drift fence Q = Fauna quadrat (fenced pit lines and, adjacent, a 200 metre x 200 metre bird quadrat) that was sampled daily for five days as described in Biological Surveys Committee of Western Australia (1984). ** The inventory prior to Appendix 1 lists the vegetation site codes and corresponding fauna sites. 51

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Page 1: VERTEBRATE FAUNA N.L. McKenzie, lK.Rolfe and W.K. …museum.wa.gov.au/sites/default/files/4. McKenzie, Rolfe, Youngson_0.pdfN.L. McKenzie, lK.Rolfe and W.K. Youngson Background The

VERTEBRATE FAUNAN.L. McKenzie, lK. Rolfe and W.K. Youngson

BackgroundThe philosophies underlying our approach to this vertebrate survey, the sampling strategy

adopted, and the actual methods we employed are described in Biological Surveys Committeeof Western Australia (1984).

Specimens representing the array of reptile and mammal species encountered in theSandstone-Sir Samuel and Laverton-Leonora Study Areas during our survey have been lodgedin the Western Australian Museum within the following series of register numbers: R62735­62872, 70827-70877, 78545-78585, 79000, R114956-1149570, 114983-114985; M20395,20396, 20404-20409, 20420, 20421, 20424, 20425, 20427, 20428, 20446-20448, 20461­20463, 20815, 20816, 20821-20823, 20825-20830, 20832-20834, 20837, 20838, 20840,

Table 2 Faunal survey of the Sandstone-Sir Samuel (Wanjarri) and Laverton-Leonora (Erlistoun) Study Areas:sampling dates and methods*.

1. Wanjarri Survey Area.

Fauna 14-21 Feb, 14-20 May, 24-29 Aug, LithologicalCode** 1979 1980 1981 Surface

1W-01 Q(M,B,R) Q(M,B,R) Q(M,B,R) Qpv/Qqz1W-02 Q(M,B,R) Q (M,B,R) Q(M,B,R) Qqc1W-03 Q(M,B,R) Q (M,B,R) Q(M,B,R) Qps/Qqz1W-04 Q(M,B,R) Q(M,B,R) Q(M,B,R) Czo+QpvlW-OS Q(M,B,R) Q (M,B,R) Q(M,B,R) Qqz1W-06 F,B,R F,M,B,R F,M,B,R Qps1W-07 F,B,R F,B,R Qps1W-08 F,B,R F,B,R Qps

2. Erlistoun Survey Area.

Fauna 22-28 Feb, 8-13 May, 18-23 Aug, LithologicalCode** 1979 1980 1981 Surface

lE-01 Q(M,B,R) Q(M,B,R) Q(M,B,R) Qpk1E-02 Q(M,B,R) Q(M,B,R) Q(M,B,R) QrmlE-03 Q(M,B,R) Q(M,B,R) Q(M,B,R) Qrd1E-04 Q(M,B,R) Q(M,B,R) Q(M,B,R) Qps/QqzlE-OS Q(M,B,R) Q(M,B,R) Q(M,B,R) QpvlE-06 F,B,R F,B,R Qps

* M = Mammal trapline (9 Elliotts, 9 Break-backs, 3 Cages)B = Bird searchesR =Reptile searchesF =Fenced pit line of six pits (4 shallow, 2 deep) with ca. 50 m drift fenceQ =Fauna quadrat (fenced pit lines and, adjacent, a 200 metre x 200 metre bird quadrat) that was

sampled daily for five days as described in Biological Surveys Committee of Western Australia(1984).

** The inventory prior to Appendix 1 lists the vegetation site codes and corresponding fauna sites.

51

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20844, 20845, 20865-20867, 20886, 20904-20910, 23501-23610, 38654-38660 and 39968.Species nomenclature in this report follows that of the Western Australian Museum.

The quadrats within the Erlistoun survey area in the Laverton-Leonora (LL) Study Area(Figures 3 and 5) allowed lithological units low in the landscape to be sampled, whereas thequadrats in the Wanjarri survey area (within Wanjarri Nature Reserve) in the Sandstone-SirSamuel (SS) Study Area were positioned to sample units high in the landscape (Figures 2 and4).

Table 1 lists the surface lithology and associated "vegetation types" known from the twoStudy Areas, while Appendix 1 indicates those that were sampled for vertebrates, cross­references the vertebrate and vegetation sample site codes, and describes the location,vegetation, floristics and substrate of each vertebrate site sampled.

Only the main vegetation types of the most extensive landform units were surveyed forvertebrates (Broad Valleys, Sal.t Lake Features, Sandplains, Dunefields and Breakaways). Thismeant, for instance, that we sampled red sandplains (surface lithology = Qps) in the contextof Broad Valleys, Dunefields as well as Sandplains (Appendix 1). It also meant that lessextensive landforms such as Granitic, Banded Ironstone and Greenstone Hills were notsurveyed at all, even though they are prominent in the Study Areas.

We surveyed the vertebrate assemblage from at least one site in five of the 13 landform.units, eight of more than 70 surface lithologies, and in 12 of over 70 vegetation types knownin the two Study Areas (Tables 1 and 2). Much of the floristic diversity of these surface-typeswas remote from our quadrats. Considering how poorly the vertebrate sampling addressed theenvironmental heterogeneity of the Study Areas, and how geographically localised ourquadrats were (Figures 2 and 3), we encountered a surprisingly high proportion of itsvertebrates (Table 3).

This coverage was achieved for passerine birds (61 %) and small ground-dwelling mammals(81 %) because most species occurred on a variety of lithologies. For instance, the "average"passerine bird occurred in a mean of 3.7 (S.D. =2.3, n =44), and an "average" small ground-

Table 3 Proportion of the Sandstone-Sir Samuel and Laverton-Leonora Study Areas' known vertebrate speciesrecorded on quadrats.

Number of SpeciesTotal On quadrats' (%)

Birds

passerinesnon-passerines

Reptilessnakeslizards

AmphibiansSmallb ground-mammals

7278"

1659c

4C

11

44 (61)19 (24)

1 (6)39 (66)

3 (75)9 (81)

* lWOl-05 (Wanjarri) & lEOl-05 (Erlistoun), excluding additional sample sites lW06-8 & lE06 (see Table 2)." Includes 30 water birds.b Indigenous species with an average adult body weight of less than 35 g (see Burbidge & McKenzie 1989).c Excluding species that only reach the periphery of the Study Areas (see reptile text).

52

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(a) WANJARRI

10

99

~ 7U~ 6(f)

LLo0::u.JID~

~ 3

2

8

(f) 7u.J

Uu.J 6a..(f)

LL 50 50::u.J 4ID~:::lZ

2

8

22 23 24 25 26 27 8 10 11 12 18 19 20 21 22

DATENo OF DAYS

FEB 1979 MAY 1980 AUG 1981

Figure 8 Species accumulation curves for small ground mammals recorded at sample sites in the a) Wanjarri(I WO I-I W06) and b) Erlistoun (I EO I-I E06) survey areas.

53

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Table 4 Species of mammals recorded in the Laverton-Leonora (E) and Sandstone-Sir Samuel (W) Study Areasindicating number of records at each vertebrate quadrat and sample site during each survey period. The threesurvey periods - Summer (February 1979), Autumn (May 19S0) and Winter (August 19SI) are indicatedin columns I, 2 and 3 respectively for each quadratlsite'.

lE' IEOI IE02 IE03 IE04 IEOS IE06LLI LL2 LL3 LL4 LLS LL6Qpk Qnn Qrd Qps/Qqz Qpv QpsS S S S S S S S S S * * S S S 0 2 S

X' X'O

Vertebrate QuadratlSiteVegetation Site'StratigraphyDrift Fence Nights

TACHYGLOSSIDAETachyglossus aculeatus

DASYURIDAEAntechinomys lanigerNingaui rideiPseudantechinus wooleyaeSminthopsis crassicaudataS. hirtipesS. macrouraS. ooldea

MACROPODIDAEMacropus robustusM. rufus

MOLOSSIDAEMormopterus planiceps'Tadarida australis

VESPERTILlONIDAEChalinolobus gouldiiEptesicus finlaysoniNyctophilus geoffroyiScotorepens balstoni

MURIDAELeporillus ?apicalisNO/omys alexisPseudomys hermannsburgensis 0 0 1Mus musculus

0102.0142.2

o 0

2. 0 0010

200

(4):1 2111

2. 0 .3.

002

S 2 1200

(2) I 0

001

:1 0 0

o 0I 1

010

020

001001

CANIDAEVulpes vulpes

FELIDAEFelis ca/us

BOVIDAE80S taurusOvis aries

CAMELlDAECamelus dromedarius

EQUIDAEEquus caballus

LEPORIDAEOryctolagus cuniculus

o 0

200

00 X'

300

X'

o X'O

x' 0 0

*

Sampling methods and dates for quadrats (IEOI-OS, IWOI-OS) and sample sites (IE06, IW06, IW07 and IWOS) areoutlined in Table 2.See Appendix l.Opportunistic records (not on quadrats).Total observations during F79 (February 1979), MSO (May 19S0) and ASI (August 19SI).These should read 10 because an additional fenced pit line was set adjacent to the bird quadrat (IE04 and IW04).

54

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lW' IWOI IW02 IW03 IW04 IW05 IW06 IW07 IW08SSI8 SS20 SS21 SS 1,2 S519 SS22 SS23Qpv/Qqz Qqc Qps/Qqz Czo+Qpv Qqz Qps Qps Qps TOTALOBS.'5 5 5 5 5 5 5 5 5 5 • 5 5 5 5 4 5 5 0 2 5 0 3 5 F79 M80 A81

----~~~

X'O 0 X' X' 0

0 0 2 () fl () () I 2 0 () ..2 () () 2 0 7 90 0 1 2 () 0 1 0 0 ..2 0 0 1 0 0 1 0 0 2 3 2 0 1 12 4 7

0 I 0 0 1 07 4 1

0 Q 2 - 1 0 0 9 22 5- ..2 I 4 5 2 4 9 12 12

() I I () 1 0 I 1 3 6

8 4 0 0 0 () 0 0 6 () 0 () () 8 10 8la 6 0 0 3. () () () 0 () 0 29 22 25

Q) 0 0 4 1 00 6 3 0 7 4

(7) 311 0 0 I 0 0 9 3 160 1 2 () 0 1 1 2

~) 0 H 9 0 13(la) 1 2 () 0 19 1 8

X' X'0 0 I 0 0 ..2 I 2 Q 2 0 - 0 1 1 4 10

1 0 0 0 2 3 I 5- 2 2 2 0 2 0 I 4(2) () 1 I 17 19 250 0 10 4 6

X' 0

2 0 0

0 ~ 0 0 ~ 010 0 0 o 30 () 10 30 0

0 X'O 0 1 0 0 X'

0 0 0 0

3 0 0

Faeces (Tachyglossus). Den (Vulpes). Warren in use (Oryctolagus). Tracks (Camelus).lS Sample includes sub-adults.( ) Sample includes post-partum, lactating or pregnant individuals.,

The form that ranges across southern Australia.Remains of old nest under breakaway.

55

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(a) WANJARRI

40

3735 3&

30 . ·3t·(j)UJU 25UJIl.(j)

LL.. 200Cl::UJIJ) 15::2;::::>z

10

DATENo OF DAYS

14 15 16 17 18 19 20 21 14 15 16 17 18 24 25 26 27 28 25

FEB1979 I MAY 1980 I AUG1981 ISEP927 12 17 18

(b) ERLlSTOUN

4038

35 35

30(j)UJU 25UJIl.(j)

LL.. 200Cl::UJIJ) 15::2;::::>z

10

5

0+--+-+......,f-+--r--+-4-+-+-+-+---1-t--t--r---r-j......,...---122 23 24 25 26 27 28 7 8 9 10 11 12 13 18 19 20 21 22 28

DATE FEB 1979 I MAY 1980 I AUG 1981 ISEP 92No OF DAYS 7 14 19 20

Figure 9 Species accumulation curves for reptiles (excluding amphibians) recorded at all sample sites(including opportunistic records) in the a) Wanjarri and b) Erlistoun survey areas.

56

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mammal in 3.4 (2.2, 9), of the eight distinct Iithological surface-types sampled (Qpk, Qrm,Qrd, Qps, Qpv, Qqc, Czo and Qqz). In contrast, the "average" lizard was recorded on only 1.8(1.1, 40) of these Iithological units; the 66% coverage for lizards may have been achievedbecause only certain lithologies (especially the widespread Qqz and Qps on which oursampling focussed) appear to have rich, compositionally distinct, lizard faunas (see Table 7).

Species accumulation curves, in relation to days of effort, were used to investigate whetherfurther sampling effort in the quadrats would have been cost-effective. Separate curves werecompiled for reptiles and small ground mammals at the Wanjarri and Erlistoun survey areas.Although the curves for survey areas are clearly beyond a final detlection point (Figures 8 and9), the reptiles of surfaces such as Qrd were clearly under-sampled (Figures 10 and 11). Theseasonally tiered pattern general to the curves indicates the importance of sampling duringseveral seasons. The results of the single day of reptile sampling in September 1992 implythat a longer session in spring would have added species. For example, if Spring records areremoved from the Kurnalpi-Kalgoorlie quadrat lists (McKenzie et at. 1992), nearly half thespecies list of ground-dwelling vertebrates is eliminated.

The analysis package PATN (Belbin 1989) was used to seek patterns of species compositionin the data matrices. The clustering techniques selected were described in McKenzie et at.(1991 b). The species assemblages from the quadrats were used as the input data. The presenceand absence of species on the quadrats, rather than their relative abundance, was used becauselimitations in sampling techniques, aggravated by staff and time limitations, precluded reliableestimates of relative abundance (Austin 1984, McKenzie et al. 1991a). Previous studies haveshown that our broad-scale sampling techniques are also unsuitable for snakes, monitors andnon-passerine birds, so they were excluded from the analyses (McKenzie 1984, McKenzie etal. 199Ia).

Table 5 Bat species recorded from the Wanjarri and Erlistoun survey areas during the survey of the Sandstone­Sir Samuel and Laverton-Leonora Study Areas.

SITE': 2 3 4 5 6 ~

I

Tadarida australis * * *Mormopterus planiceps * * + *Chalinolobus gouldii * * * * * *Scotorepens balstoni * * * * * *Eptesicus finlaysoni * * *Nyctophilus geoffroyi * *

8

**

*

, I = 27"17'5 120"40'E (granite outcrop);2 =27°20'5 120"40'E (Bullock Well, Mulga);3 =27"32'5 120"43'E (Dingo Pool, Mulga);4 =27"21'5 120"36'E (breakaway pool at 1W04);5 27"24'5 120"41 'E (red sand dune near IW06);6 27"24'5 1200 39'E (Homestead Dam, Mulga).7 =28"16'5 122"06'E (lake Ikm north of 1E03).8 =28°22'5 I 22"OTE (dam, Mulga).

+ =recorded on 25 September 1992 from sonar signature using ANA BAT 11 (Corben 1991).

57

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Qqc(1W02)

20

" 17I.~ 14

U~ 12</)

~ 10

a::l:i 8::;::0Z

44

DATENo OF DAYS

14 15 16 17 18 19 20 21

FEB 1979

14 15 16 17 18 24 25 26 27 28 25

MAY 1980 I AUG 1981 I SEP 9212 17 18

CZQ & Qpv (1W04)20

</) ,.UJUUJ 12a. 12 12</)

u. 100a::UJ 8<D::;::0Z

18,.

DATENo OF DAYS

14 15 16 17 18 19 20 21 14 15 16 17 18 24 25 26 27 28 25

FEB 1979 I MAY 1980 I AUG 1981 ISEP 927 12 17 18

Qpv (1W01)20

</)

"UJUUJ 12a.</)

u. 10 '90a::UJ 8<D::;::0Z

18

"

DATENo OF DAYS

14 15 16 17 18 19 20 21 14 15 16 17 18 24 25 26 27 28 25

FE81979 I MAY 1980 I AUG 1981 ISEP 927 12 17 18

58

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Qps (1W03 & 1W06)

'018

U)UJ(3g: 12U)

0 10

'"1iS 8::;;::JZ

20

10

14 15 16 17 16 19 20 21

FEB 19197 !

14 i5 16 17 18 24 25 26 27 28 25

MAY 1980 12 \ AUG 1981 171 s~: 92

Species accumulation curves for reptiles (excluding amphibians) recorded on each surface sampledin the Wanjarri survey area. Accumulation curves for the individual sample sites on each surfaceare also included for comparison.

14 i5 16 17 18 24 25 26 27 28 25

MAY 1960 121 AUG 1981 171 s;; 92FEB 1979

14 15 16 17 18 19 20 21

Qqz (1W05)

20

18

"U)

" I Qqz IUJ(3 ---1W05UJ 12"-1/)

10 10 10"- 100

'"UJ1lJJ

::;;::JZ

4 54

DATENo OF DAYS

..~ Figure 10

59

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Qpv (1E05)2<J

",.(f) ,.wu

1 Qpv fw 12a.. . --lEOS(f)

u. 100

'"W •"'::; 7

'" 6z

8 9 10 11 12 13 18 19 20 21 22 28

MAY 1980 I AUG 1981 I SEP 9214 19 20

FEB 1979

oL4=~~:re::::22232<4252627287

DATENoOF DAYS

Qrd (1E03)20,.

2324252627287

,.~ I.U~ 12(f)

~ 10

'":,g •::;

'"Z

DATENo OF DAYS

4

FEB 1979

17

I Qrd I. ---1E03

9

8 9 10 11 12 13 16 19 20 21 22 28

MAY 1980 I AUG 1981 I SEP 9214 19 20

Qps (1 E04)

8 9 10 11 12 13 18 19 20 21 22 28

MAY 1980 I AUG 1981 I SEP 9214 19 20

FEB 1979

2324252627287

2.

22

20,.(f)w ,.Uwa.. \4(f)

u. 120

'"11

w 10

"'::;

'"z

DATENo OF DAYS

60

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Qrm (1E02)

20

18

16

~~--1E02

DATENoOF DAYS

FEB 1979

8 9 10 11 12 13 18 19 20 21 22 28

MAY 1980 141 AUG 1981 19 i Si: 92

Qpk (1E01)

20

18

16

if) 14W(3 I Qpk Iw 12a. --1E01if)

"- 100

'" 9w 8co::;;::JZ

DATENo OF DAYS

22 23 24 25 2f5 27 28 7

FE8197971

8 9 10 11 12 13 18 19 20 21 22 28

MAY 1980 I AUG 1981 ! SEP 9214 19 I 20

A~ Figure 11 Species accumulation curves for reptiles (excluding amphibians) recorded on each surface sampled

in the Erlistoun survey area. Accumulation curves for the individual quadrats on each surface arealso included for comparison.

61

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Table 6 Data matrix of small ground-dwelling mammals re-ordered according to UPGMA classification outputs(Belbin 1989). The quadrats were classified according to similarities in their species composition, andthe species according to their co-occurrence at the same quadrats and sample sites.

QuadratlSite Codes!

SpeciesPseudantechinus wooleyae

Sminthopsis hirtipes

Notomys alexisNingaui ridei

Antechinomys laniger

Pseuomys hermannsburgensisSminthopsis ooldea

Sminthopsis macroura

Mus musculusSminthopsis crassicaudata

Lithological origin2

! Quadrat & sample site codes are printed vertically.Q = Quaternary; T = Tertiary.

I I 11 I I1I 1I 1Ie e e e wwww ew ew0000 0000 00 002135 1253 44 66

** *

* * * ** * * * ** * ** * * *

* * * * * * * * * * ** * ** **

* * * * * *

* * * * * ** * *QQQQ QQQT TT TT

MammalsEighteen species of native mammal were extant on the quadrats during the survey (Table

4), These comprised two species of kangaroo, 9 small ground-dwelling mammal species (7dasyurids and 2 rodents), the Echidna, and six species of insectivorous bat Eight species ofintroduced mammal were also encountered (including Mus musculus). In addition, more than40 Goats (Capra hircus) were seen in October 1985 (CALM files), and several herds of 20 ormore were seen by the authors in September 1992.

According to Western Australian Museum records, 15 indigenous and two introducedmammals were previously known from the Study Areas. Two of the indigenous species werenot recorded during our field work: Macrotis lagotis (M632, Laverton, 1924) andMyrmecobius fasciatus (M351, Ida Hill Mine, 1918, 28"41 'S 122"24'E); both are known tohave become extinct or very much rarer throughout most of their former range since Europeansettlement (Burbidge and McKenzie 1989).

Another 14 native species have been identified from bone material in exposed, superficialsub-fossil deposits in the Youanrni-Leonora Study Area (Figure 1), immediately to the southof the Study Areas described herein (Appendix 13 in Henry-Hall 1990). Twelve of thesespecies are known to have become extinct or very much rarer throughout mainland Australiasince European settlement (Burbidge and McKenzie 1989): Dasycercus cristicauda,

62

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9

8

(a)

7

6

5

10

3

4

2

8

9

6

7

(b)

5 4

10

11

3

2

Figure 12 Three-dimensional ordinations of the small, ground-dwelling mammal data from sites at Wanjarri(lWOI-lW06) and Erlistoun (lEOI-IE06). Minimum Spanning Trees have been superimposed.Shaded wedges represent projections into the 3rd dimension. (a) Species ordinated according totheir co-occurrences at the same sites. Stress=0.037. (b) Sites ordinated according to similarities intheir species composition. Stress=0.07. See text for details of analyses.

a) Species ordination.

KEY:

I. Sminthopsis crassicaudata2. Mus musculus3. Sminthopsis macroura4. Sminthopsis ooldea5. Pseudomys hermannsburgensis6. Antechinomys laniger7. Ningaui ridei8. Notomys alexis9. Sminthopsis hirtipes10. Pseudantechinus wooleyae

b) Site ordination.

KEY:

Red Tertiary sand. hi~h in the landscape:1. lE06 & 1W06: as dunes (Qps) at the top of the landscape.2. lW03: as sheets mantling Quaternary soil plains (QpslQqz).

Quaternary surfaces at middle levels in the landscape:3. lWOS: Soil plain (Qqz).4. lW02: Gibber slope (Qqc).5. lW01: Alluvium (Qpv) in freshwater drainage line incising the Qqz.

Quaternary surfaces lower in the landscape6. lEOS: Alluvium (Qpv) in freshwater drainage line incising a QpslQqz

surface.7. lE03: Brownish sand (Qrd) peripheral to Qnn.8. lE02: Qccluded drainage of saline alluvium (Qnn) at the bottom of

the landscape.9. lEOl: Gypsum dune (Qpk) peripheral to Qnn.

Tertiary surfaces. medium to low in the landscape10. lE04: Red sand sheet, mantling Quaternary soil plains (QpslQqz).11. 1W04: As duricrust breakaway complex (Czo) and associated

Quaternary alluvial deposits (Qpv).

63

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Table 7 Species of reptiles recorded in the Laverton-Leonora (E) and Sandstone-Sir Samuel (W) Study Areasindicating number of records at each vertebrate quadrat and sample site during each survey period. Thethree survey periods - Summer (February 1979), Autumn (May 1980) and Winter (August 1981) - areindicated in columns 1,2 and 3 respectively for each quadratlsite'.

Vertebrate QuadratlSiteVegetation Site2

StratigraphyDrift Fence Nights

Cye/orana mainiCye/orana platycephalaNeobatrachus ?kunapalari

Diplodactylus conspicillatusD. elderiD. pulcherD. squarrosusD. strophurusD. wellingtonaeGehyra purpurascensG. variegataHeteronotia binoeiUnderwoodisaurus miliiRhynchoedura ornata

lE'

040200150

100

100100010001o 223113100101

IEOI IE02 IE03 IE04 IEOS IE06LLl LL2 LL3 LL4 LLS LL6Qpk Qrm Qrd Qps/Qqz Qpv Qps555 555 5 5 5 5 * * 5 5 5 o 2 5

0 0

0 0 2 0 0

0 0S

100

SS S 0 0

Delma butleriD. nasutaLialis burtonis 100

Soo

oo 3 0

Caimanops amphiboluroidesCtenophorus caudicinctusC. fordi 2 0 3C. inermis 1 1 0 0 I 0C. isolepis 2 3 2 2 2 0 - I 0C. reticulatus 2 0 0C. salinarum 1 0 2 0 0 I 2 I 2C. scutulatus 1 0 0 0 0Moloch horridus 1 0 2Pogona minor 0 0 2 0 0 I 0 I S

, Sampling methods and dates for quadrats (1E01-05, IWOI-05) and sample sites (lE06, IW06, IWO? andIW08) are outlined in Table 2.

2 See Appendix I.) Opportunistic records (not on quadrats).4 Total observations during F79 (February 1979), M80 (May 1980) and A81 (August 1981); excludes records

from October 1987 and September 1992 visits.* These should read 10 because an additional fenced pit line was set adjacent to the bird quadrat (I E04 and

IW04).S Recorded in September 1992.

64

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lW' lWOl lW02 lW03 lW04 lW05 IW06 lW07 lWOSSSlS SS20 SS2l SSl,2 SSl9 SS22 SS23Qpv/Qqz Qqc Qps/Qqz Czo+Qpv Qqz Qps Qps Qps TOTALOBS"

5 5 5 5 5 5 5 5 5 5 * 5 5 5 5 4 5 5 0 2 5 0 3 5 F79 MSO ASl

I 0 0 2 0 0 I 0 0 5 4 0I 0 0 0 0 4 0 05 0 0 7 0 0 4 0 0 0 0 0 I 0 2 0 20 9 0

0 0 0 0 3 0 I 7 0S

0 0 2 0 0 3 0 02 0 0

- 0 1 I 0 I0 0 1 0 0 0 0 0 0 4 I 0

0 0 II 0 S 7 0 1 8 3 250 I 2 0 0 4 I 4

I 0 00 0 4 0 0 6 0 1

1 0 0 I 1 00 0 1 - 0 1 0 4 2

I 0 0

I 0 0 0 1 0 I I 00 0 2 1 0 2 I 0 4

2 0 36 0 0 1 0 0 I 0 0 0 0 10 2 02 0 0 3 0 0 2 0 0 0 1 II 6 32 0 0 0 5 0 II 0 0 4 I 5

2 0 0I 0 0 2 0 2

2 0 0 3 0 3

65

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Table 7 (conI.)

Vertebrate QuadratfSiteVegetation Site2

StratigraphyDrift Fence Nights

CryplOblepharus carnabyiC plagiocephalusCtenotus ariadnaeC calurusC grandisC greeriC helenaeC leonhardiiC pantherinusC quattuordecimlineatusC schomburgkiiEgernia depressaE.jormosaEremiascincus richardsoniiLerista bipesL. deserlOrumL. muelleriMenetia greyiiMorethia butleriTiliqua multijasciataT. occipitalis

Varanus brevicaudaV. caudolineatusV. gouldiiV. panoptes

Furina ornataPseudechis australisPseudonaja nuchalisRamphotyphlops hamatusVermicella bertholdi

Species Richness

lE'

120

120

001004002002

o 1

010100

1 0o 0

s33

IEOILLlQpk555

S

So 0

S

7

IE02LL2Qrm555

100

100

3

IE03LL3Qrd555

S

S

S

100

9

1E04LL4Qps/Qqz

5 * *

200120

3 2

o 2

100

100

12

IE05LL5Qpv555

S

001

8

IE06LL6Qps025

2

I Sampling methods and dates for quadrats (1E0 1-05, IWO 1-05) and sample sites (I E06, IW06, IW07 andIW08) are outlined in Table 2.

2 See Appendix I., Opportunistic records (not on quadrats).4 Total observations during F79 (February 1979), M80 (May 1980) and A81 (August 1981); excludes records

from October 1987 and September 1992 visits.* These should read 10 because an additional fenced pit line was set adjacent to the bird quadrat (lE04 and

IW04).S Recorded in September 1992.

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lW' IWOI IW02 IW03 IW04 IW05 IW06 IW07 IWOS551S 5520 5521 551,2 5519 5522 5523Qpv/Qqz Qqc Qps/Qqz Czo+QpvQqz Qps Qps Qps TOTALOB5'

5 5 5 5 5 5 5 5 5 5 * 5 5 5 5 4 5 5 0 2 5 0 3 5 F79 MSO ASI

2 0S

3 0 3 I 00 0 0 I 0

0 0 I 0 02 0 0

0 0 I 2 0 0 4 4 I2 0 0 5 0 0 I 0 0 2 0 0 10 0 0

0 I I 0 3 3 2I 0 0 0 2 I 0 2

I 0 0 5 0 I 0 0 2 0 0 2 I 0 11 3 22 0 0 0 0 3 0 0

2 0 0 2 0 0I 0 0 1 0 0

5 I 0 0 1 0 00 0 1

5 5 1 0 5I 1 I 0 0 3 0 0 5 1 30 I 0 I 3

I 0 0 2 0 0I 0 I

1 0 02 0 0 I 0 2 3 I 3

I 0 0I 0 0 I 0 0

1 I 00 1 0

1 0 0 1 0 00 0 2 0 0

S

23 10 7 II 14 6 6 2 5 57

67

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Phascogale ca Iura, Chaeropus ecaudatus, Isoodon auratus, Trichosurus vulpecula,Onychogalea lunata, Leporillus apicalis, L. conditor, Notomys amplus, N. longicaudatus,Pseudomys fieldi and Rattus tunneyi. Except for the Dasycercus, there is no evidence thatthey persist in the district. CALM Departmental files include a letter from J.L. Bannister,dated 12 December 1969, noting that T.K. Moriarty had sent a female Dasycercuscristicaudata to the W.A. Naturalists Clubs Wildlife Show in 1967, but the specimen wassubsequently lost. At that time, Bannister was the Curator of Mammals at the WesternAustralian Museum and Moriarty owned and operated the Kathleen Valley pastoral lease (=Wanjarri). G. Connell (pers. comm.) trapped D. cristicaudata in the south-western part of theWanjarri Nature Reserve in 1991.

Bones of Pseudomys holami and P. desertor were also found in the sub-fossil deposits fromthe Youanmi-Leonora Study Area. However, P. holami was not trapped during our survey, orin other Study Areas in the northern Goldfields even though appropriate habitats weresampled (Dell and How 1992, How and Dell 1992). Noting that the species was readilytrapped in survey areas to the south (e.g. Dell and How 1988), the sub-fossil record from theYouanmi-Leonora Study Area may represent the northern limit of this species' range. P.desertor may still be extant in the Study Areas.

60

50

Cl)UJ 40()UJa.Cl)

L.l. 3000::UJco::lE

20:::>z

10

0

~•..

I-.197901980.1981

I- r'

I- f· f-.

---- . ... ...

f-· ..-

.d2 3 4 5 6 7 8 9 10 11 12 13 14 15

NUMBER OF DAYS

Figure 13 Species accumulation histogram for reptiles (excluding amphibians), showing seasonal differencesin the number of species encountered per day of sampling effort. Results from the Wanjarri andErlistoun survey areas are combined.

68

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The extant native mammal fauna comprises only Eyrean species. There arc several speciesin common with the adjacent deserts (the Helms and Keartland Botanical Districts): e.g.Sminthopsis ooldea. S. hirtipes. Ningaui ridei. Notomys alexis and Pseudomyshermannsburgensis. This retlects the mozaic of red sandplain and sand dune surfaces scatteredthrough the northern Goldfields. The presence of Pseudantechillus wooleyae and Eptesicusjinlaysolli, rather than their desert counterparts (Pseudantechinus macdonnellellsis andEptesicus baverstocki occur in the Helms District), is consistent with the phytogeographicposition of our Study Areas in the Austin Botanical District, as discussed in Keighery et al.(this publication).

Bats were sampled using mist nets, a bat trap and spotlight shooting. Sites were usually on

Table 8 Amphibians and reptiles from the Sandstone-Sir Samuel Study Area not recorded during the survey.~-------_._-_.------_.._._--_.__._---

Species

Limnodynastes spenceriPseudophryne occidenlalis

Diplodactylus elderiD. squarrosusNephrurus verrebralisN. wheeleriOedura marmorata

Pygopus nigriceps

Ctenophorus ornatusC. scutulatusTvmpanocryptus cephala

Cryptob!epharus plagiocephalusEgernia inornalaE. kintoreiE. striataLerisla deserrorumTiliqua occipitalis

Varanus eremiusV. giganteusV. gouldii

Morelia stimsoniAcanrhopis pyrrhusDenisonia fasciataPseudechis australisP. burleriPseudonaja modestaRhinoplocephalus monachusVermicella bertholdiV. semirascialaRamphotyphlops waitti

* WAM Western Australian Museum, Perth

Specimen*

WAM R 104473WAM R 100534

WAM R 26878WAM R 104471WAM R 48031WAM R 8942WAM R 84357

WAM R 1103

WAM R48026WAM R 27219WAM R 100967

WAM R 82561WAM R 48037WAM R 27228WAM R 100964WAM R 19766WAM R 37786

WAM R 27230WAM R 27232WAM R 12406

WAMR 1417WAM R 30599WAM R 40522WAM R 12599WAM R 7627WAM R 1122WAM R 19777WAM R 31671WAM R 40523WAMR 19778

69

Source Location

YakabindieBarr Smith Range

Montague RangeYakabindieWanjarriSandstoneAnketell Outstation

Booylgoo Spring

WanjarriWanjarriWanjarri

Yeelirrie HomesteadWanjarriWanjarriWanjarriWanjarriWanjarri

WanjarriWanjarriWanjarri

Booylgoo SpringAlbion DownsWanjarriWanjarriBooylgoo SpringBooylgoo SpringWanjarriWanjarriWanjarriWanjarri

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pools, places where bats congregate to drink. The observed assemblages are listed in Table 5.A solitary N. geoffroyi was found roosting in a Mulga spout during the day.

Tadarida australis seldom foraged less than 10 metres above the ground, and always wellclear of the vegetation canopy. It was often recorded in pairs. Mormopterus planiceps andChalinolobus gouldii foraged just above the open woodland canopy (6 9 metres above theground), although the C. gouldii flew more slowly and often foraged between tree canopies.Scotorepens balstoni and Eptesicus jinlaysoni were seen foraging the gaps between trees andshrubs, although E. jinlaysoni flew more slowly and closer to the vegetation, with frequentchanges in direction. Two Nyctophilus geoffroyi were tracked at night using luminescentCyalume beads (see Buchler 1976) in a low open Mulga (Acacia aneura) woodland near theWanjarri homestead. They fluttered slowly along less than two metres above the ground, oftenchanging direction and occasionally diving almost vertically to its surface. These differencesin bat foraging micro-habitats are consistent with wing-shape differences described for thesespecies by Fullard et al. (1991) and McKenzie and Start (1989).

Data on small ground mammals collected during the trapping program are summarised inTable 4. The trap-effort values at the top of this table provide a relative measure of samplingeffort because all small ground mammals recorded on quadrats were trapped.

The results of the numerical classifications are presented as a re-ordered data matrix inTable 6. To confirm the robustness of the clusters, species and quadrats were ordinated in 3­dimensional space using Semi-strong Hybrid Scaling (Belbin 1991), then a MinimumSpanning Tree was superimposed (Figure 12). These analyses revealed a gradient in thecomposition of the small ground mammal assemblages in the survey areas that could beexplained in terms of surface lithology.

The upper levels in the landscape are Tertiary surfaces. These were characterised byParantechinus wooleyae where duricrust is exposed as breakaways along eroding edges of theuplands, and by Smimhopsis hirtipes on the red sand dune systems (Figure 12 and Table 6).The Tertiary surfaces also supported Notomys alexis, Ningaui ridei, Sminthopsis ooldea andPseudomys hermannsburgensis, but these four species also occur down the landscape sequenceto levels where sheets of Tertiary sands have mantled, or are mixed into the upper profile, ofheavier Quaternary surfaces. The last two of these "Tertiary" species extend into Quaternarysequences, although Sminthopsis ooldea avoids the heavier soils.

The lower levels in the landscape are Quaternary surfaces. Antechinomys laniger favouredthe heavy soils of Quaternary plains, away from the salt lake systems. Sminthopsis macrourawas found on all Quaternary surfaces except the salt lake bottoms and gypsum dunes. At thebottom of the landscape, the salt lake surfaces and their peripheral features supported the S.crassicaudata and Mus musculus populations. A similar pattern in composition was noted inthe Kurnalpi-Kalgoorlie Study Area, although results from Kurnalpi indicate that S.crassicaudata can disperse across Tertiary surfaces (McKenzie et al. 1992).

Although a session of sampling in spring would have captured additional species onquadrats, the levels of sampling were sufficient to show that compositional gradients reflectedthe geomorphological sequences in the landscape's surface stratigraphy (see Figure 12).

Reptiles and AmphibiansThree frogs and 54 reptile species were recorded in the Sandstone-Sir Samuel and

70

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Table 9 Amphibians and reptiles from the Laverton-Leonora Area not recorded during the survey.

Species

Liml1odYl1astes spel1ceriPseudophryl1e occidel1talis

Diplodactylus assimilisD. gral1ariel1sisD. pulcherD. stel10daclylusNephrurus le visN vertebralis

Pygopus l1igriceps

Ctel1ophorus caudicil1ctusTvmpal10cryptus cephala

Ctel10tus ariadl1aeC. calurusC gral1dis

C hal1lol1iC leol1hardiiC pial1kaiC qualluordecimlil1eatusC severusEgernia depressaEformosaE il10rnataE richardiE striataEremiascil1cus richardsol1iiLerista bipes

Varal1us eremiusV pal10ptesV tristis

Morelia stimsol1iDemal1sia psamophisDel1isol1ia fasciataPseudechis bUileriPseudol1aja modestaP. l1uchalisRhil1oplocephalus mOl1achusVermicella bertholdiV fasciolataV semi/asciataRamphotyphlops el1doterusR. waitii

, WAM Western Australian Museum, Perth

Specimen'

WAM R 1248WAM R 21536

WAM R 20651WAM R 97289WAM R 1246WAM R 85633WAM R 85634WAM R 78156

WAM R 85592

WAM R 25946WAM R 1240

WAM R 85714WAM R 85781WAM R 86217WAM R 86453WAM R 1234WAM R 85690WAM R 89073WAM R 20617WAM R 1200WAM R 20659WAM R 86081WAM R 25945WAM R 86136WAM R 17897WAM R 85884

WAM R 11609WAM R 98075WAM R 100396

WAM R 13852WAM R 87884WAM R 1426WAM R 22395WAM R 15052WAMR11047WAM R 11475WAM R 85353WAM R 1751WAMR 13854WAM R 13855WAM R 21336

7\

Source Location

LavertonWhite Cliffs

White Cliffs60km SE LeinsterLavertonPt Salvation39km E Laverton10km S Leonora

39km E Laverton

40km NE LavertonLaverton

39km E Laverton39km E Laverton39km E LavertonLavertonLaverton39km E Laverton39km E LavertonIlkm E LeonoraLavertonWhite Cliffs39km E Laverton40km NE Laverton39km E LavertonCosmo Newbery39km E Laverton

LavertonBundarra Station39km E Laverton

Cosmo NewberyCosmo NewberyLavertonLavertonCosmo NewberyLeonoraLavertonWhite Cliffs StationLavertonCosmo NewbervCosmo NewberyLaverton

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5b

10

3

5a

Figure 14 Three-dimensional ordination of the reptile data (except snakes and moniters) from sites atWanjarri (lWOI-IW08) and Erlistoun (lEOI-IE06). Stress=0.14. Sites are ordinated according tosimilarities in their species composition. A Minimum Spanning Tree has been superimposed.Shaded wedges represent projections into the 3rd dimension. See text for details of analyses.

KEY:Ouaternary soils low in the landscapeI. lE02: Occluded drainage of saline alluvium (Qrm).2. lE01: Gypsum dune (Qpk) peripheral to Qrm.3. lE05: Alluvium in freshwater drainage line (Qpv) incising a QpslQqz surface.4. lE03: Brownish sand (Qrd) peripheral to Qrm.

Red Tertiary sandSa-d. lW06, lWOS, lE06 & lW07 (respectively): as dunes at the top of the landscape (Qps).6a-b. lE04 & lW03 (respectively): as sheets mantling Quaternary soil plains (QpslQqz).

Ouaternary soils at middle levels in the landscave7. lW05: Soil plain (Qqz).8. lW02: Gibber slope (Qqc).9. lW01: Alluvium (Qpv) in freshwater drainage line incising soil plain (Qqz).

Tertiary breakaway complex10. lW04: Duricrust outcrop (Czo) and associated Quaternary alluvial deposits (Qpv).

72

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Laverton-Leonora Study Areas during our survey. The reptile species comprised 11 geckos, 3legless lizards, 10 dragons, 21 skinks, 4 goannas and 5 snakes. These are listed in Table 7.

A computer search of the W.A. Museum collection revealed records of another 2 amphibianand 31 reptilian species from the Study Areas during the period of European settlement(Tables 8 and 9). Thus, a minimum of 5 amphibian and 85 reptile species were extant in theseStudy Areas at the time of European settlement: 78 reptiles in the Laverton-L,e(mora and 64in the Sandstone-Sir Samuel Study Area.

However, one of the amphibians and 10 of the lizards listed in Tables 8 and 9 are onlyknown from the periphery of our Study Areas; their geographic ranges are centred elsewhere(Pseudophryne occidentalis, Diplodactvlus assimilis, D. granariensis, D. stenodactvlus,Nephrurus ICl'is, N. Ivheeleri, Oedura marmorata, Ctenotus piankai, C. severtls, Egerniakintorei and E'. richardi).

The proportion of the known reptile species richness that we recorded from quadrats islisted in Table 10. Clearly, our survey methods were less effective for monitors and snakes.Similar biases in reptile sampling have been discussed by McKenzie et al. (1987).

The rich reptile lists from the Sandstone··Sir Samuel and the Laverton-Leonora StudyAreas reflects long-term collecting by Eric Pianka (Pianka 1969, 1989) at sites in the easternportion of our Study Areas, and by T.K. Moriarty at Wanjarri. Their richness compares withthe 66 species known from the Kurnalpi-Kalgoorlie Study Area to the south (McKenzie et al.1992).

Although they have not actually been recorded within the Study Areas, at least another 2amphibian and 5 lizard species might be expected on the basis of their known distributions inWestern Australia: Cyclorana australis, Neobatrachus wilsmorei, Gemmatophora longirostris,Ctenows uber, C. brooksi, C. colletti and C. dux.

Aside from wide-ranging species, there are three more specialised components in the reptilefauna:(I) Species with distributions centred on the heavier soils of the arid pastoral lands to thenorth, south and west (e.g. Diplodactylus squarrosus, D. wellingtonae, Caimanopsamphiboluroides, Ctenophortls salinarum, C. scutulatus, Ctenotus severus, Egernia depressa,E. formosa and Tiliqua occipitalis).

Table 10 Proportion of the Sandstone-Sir Samuel and Laverton-Leonora Study Areas' known reptile speciesrichness recorded on quadrats, by family.

Known l

12 geckoes4 legless lizards

12 dragons24 skinks

7 monitors16 snakes

75 TOTAL

From Quadrats2

10 (83%)2 (50%)9 (75%)

17 (71%)3 (43%)1(6%)

42 (56%)

From both Study Areas, excluding peripheral species (see Tables 8 & 9).

Excludes the additional sample sites IW06-8 & IE06 (see Tables 2 & 7)

73

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(2) Species that favour the red sandplains and dunefields that dominate the Great VictoriaDesert to the east, and are scattered across the two Study Areas (e.g. Ctenophorus isolepis, C.fordi, Ctenotus calurus, C. quattuordecimlineatus, Lerista bipes and L. desertorum).(3) A Bassian component with distributions centred on semi-arid districts, that comprisesthe amphibian Pseudophryne occidentalis and the lizards Diplodactylus granariensis,Phyllurus milii, Ctenophorus omatus and Egemia richardi. None of these extend into thewettest areas of mesic South-western Australia.

The dominance of arid zone reptiles is consistent with phytogeographic patterns; the StudyAreas are entirely within the Eremaean Phytogeographic Province. They are in the north­eastern (inland) corner of the Austin Botanical District, and remote from the South-WesternInterzone (Beard 1980).

A more prolonged session of sampling in the springtime would have captured additionallizard species on quadrats; even a single day of sampling in the spring of 1992 revealedadditional species at six quadrats (Table 7, Figures 9-11). Quadrats on equivalent surfaces inadjacent Study Areas were richer in species (Dell and How 1992, How and Dell 1992).Nevertheless, from Figure 13 it is clear that the summer sampling session was more thantwice as effective as the winter and autumn visits.

The richest quadrat for reptiles was 1W04, an ecotone between Czo (Tertiary laterite) andQpv (a gully incising the plateau edge - see Appendix 1). The lizard assemblages at two ofthe quadrats on alluvial surfaces OWOS =Qqz, lW02 =Qpc) were clearly incomplete (Figure10). Their richness may have been reduced by over-stocking; the Mulga tree canopy on thesequadrats was dead and the surface of the soil was bare and sheet-eroded (see Appendix 1).

Figure 14 presents a 3-dimensional scatterplot derived by ordinating the quadrats accordingto similarities in their lizard species composition, then superimposing a Minimum SpanningTree (the analysis pathway described earlier in this paper). These numerical analyses reveal astrong relationship between the composition of lizard assemblages and surface lithology. Thelevels of sampling were sufficient to show that gradients in composition reflected thegeomorphic sequences in surface stratigraphy across the landscape. Analysis of data fromanother Eastern Goldfields Study Area yielded a similar conclusion (McKenzie 1984). Incontrast, rainfall and temperature attributes are usually found to be significant as scalars oflizard composition in study areas with significant climatic gradients, such as areas withgreater relief or those that cover entire districts (e.g. McKenzie et al. 1987, How et al. 1987,Gambold and Woinarski in press).

BirdsNinety-three species of bird were recorded in the Sandstone-Sir Samuel and Laverton­

Leonora Study Areas during our survey. Sixty-three of these (Table 11) were recorded onsurface-types consistently sampled by both quadrat and opportunistic techniques. Speciesknown from the Sandstone-Sir Samuel and Laverton-Leonora Study Areas, but not recordedon quadrats during our surveys, are listed in Table 12.

Additional birds have been recorded on adjacent salt lakes to the north of the Sandstone-SirSamuel Study Area (Curry 1979, Storr 1985) and to the east of the Laverton-Leonora StudyArea (Storr 1986). Species recorded on Lake Violet (26°39'S 120°16'E) were: Blue-billedDuck, Eastern Golden Plover, Greenshank, Sanderling, Red-necked Stint, Long-toed Stint,

74

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Pectoral Sandpiper, Curlew Sandpiper, Oriental Pratincole and Silver Gull. Three other birdsrecorded in adjacent areas to the north and cast of both Study Areas were the SamphireThornbill, Calamanthus and Painted Finch (Storr 1985, 1986).

Thirty-two of the 61 non-passerines listed in Table 12 arc swamp or lake birds; no quadratswere sampled in these habitats during our field work. Many of the other non-passerines in thistable arc species near the edge of their geographic range (e.g. Regent Parrot, Purple-crownedLorikeet and Elegant Parrot); others have declined since settlement (Malleefowl, PrincessParrot), or arc vagrants or seasonal migrants (e.g. Black-cared Cuckoo, Rainbow Bee-cater).Of the 27 passerine species that wc failed to record on quadrats, the Grey-crowned Babblerand Purple-gaped Honeyeater (further records arc needed to confirm this record) arc species atthe limits of their range, species such as the Black Honeyeater, Grey Honeyeater and White­browed Woodswallow arc vagrants or seasonal nomads, and the Western Wedgebill is nowscarce through much of its former range. The White-plumed Honeyeater, the Orange Chatand the White-fronted Chat, Rufous-crowned Emu-Wren, Striated Grasswren and Spinifex­bird favour habitats that wc did not sample using bird quadrats.

The 151 species recorded since European settlement in the Sandstone-Sir Samuel andLaverton-Leonora Study Areas comprises 80 non-passerines and 71 passerines. The Eyreancomponent is dominant because of the geographical location of the Study Areas and theirvariety of distinctly Eremaean vegetations.

*GEOLOGY *QUADRAT SIMILARITY COEFFICIENT (CZEKANOWSKI)

0.16 0.29 0.42 0.55 0.68 0.81

Opk 1E01

Onn 1E02

Ord 1E03

Ope 1W02

Opv 1E05

Czo+Opv 1W04

Opv/Oqz 1W01

Oqz 1W05

Ops/Opz 1E04

Ops/Oqz 1W03

* SEE TABLE 2

I SALINE

I

I

I ALLUVIAL

I RED SAND

Figure 15 Bird quadrats classified according to similarities in their passerine species composition (excludingnon-passerines and corvids) using the Czekanowski (1932) similarity algorithm and unweightedpair-group mean averaging (U.P.G.M.A.) as a sorting strategy (Belbin 1987).

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Table 11 Species of birds recorded in the Laverton-Leonora (E) and Sandstone-Sir Samuel (W) Study Areasindicating number of sightings at each vertebrate quadrat during each survey period. The threesurvey periods - February 1979 (Summer), May 1980 (Autumn) and August 1981 (Winter) areindicated in columns I, 2 and 3 respectively for each quadrati.

Stratigraphy Qpk Qrm Qrd Qps/Qqz QpvVegetation Site2 LLl LL2 LL3 LL4 LL5Vertebrate Quadrat IEOI lE02 lE03 IE04 IE05

NON-PASSERINESEmu T 2 -Hoary-headed GrebeLittle Pied CormorantWhite-faced HeronPacific HeronMountain DuckGrey TealBlack DuckPink-eared DuckWood DuckCollared SparrowhawkLittle EagleWedge-tailed Eagle 8 - - .J. - 1 -Spotted HarrierBrown Falcon - 1Australian Kestrel - I - - .2 1Australian HobbyBlack-tailed Native-henEurasian CootAustralian BustardBanded PloverBlack-fronted PloverRed-capped PloverCommon SandpiperBlack-winged StiltBush Stone-curlewDiamond Dove 8Common Bronzewing 4 -Crested Pigeon - 2 5 4Mulga Parrot 4 5

Each quadrat was sampled for five days over three seasons.See Appendix I.Opportunistic sightings (not on quadrats).Total observations during F79 (February 1979), M80 (May 1980) and A8I (August 1981).

T Tracksx No season assigned for opportunistic sightings..2 All sightings of birds in flight.* Recorded in August 1982.99 Unspecified number of individuals (99+).

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Table 11 (cont)

StratigraphyVegetation Site2

Vertebrate Quadrat

Ring-necked ParrotBourke's ParrotBudgerigarCockatielGalahPallid CuckooHorsfield's Bronze-CuckooTawny FrogmouthAustralian Owlet-nightjarRed-backed Kingfisher

QpkLLlIEOI

2

2.lQlQ1 6 2

QrmLL2IE02

Qrd Qps/Qqz QpvLL3 LL4 LL5IE03 IE04 IE05

- 3 1 2 23 2 I

~ - ~ 1 I12. I 2.1 :l 20 :l

PASSERINESWhite-backed SwallowTree MartinRichard's PipitGround Cuckoo-shrikeBlack-faced Cuckoo-shrikeWhite-winged TrillerJacky WinterHooded RobinRed-capped RobinRufous WhistlerGrey Shrike-thrushCrested BellbirdWillie WagtailCinnamon Quail-thrushWhite-browed BabblerSouthern WhitefaceWeebillBroad-tailed ThornbillYellow-rumped ThornbillSlaty-backed ThornbillChestnut-rumped ThornbillStriated Grasswren

2

- 2

3.1 ­

2

- 3 2

IlQ - I

- 1- 2 1- 5 7

6 1 10- 3

- - 3

- 1

- 2- 1

- 4 4

- 3

*

Tx1

Each quadrat was sampled for five days over three seasons.See Appendix I.Opportunistic sightings (not on quadrats).Total observations during F79 (February 1979), M80 (May 1980) and A81 (August 1981).TracksNo season assigned for opportunistic sightings.All sightings of birds in flight.Recorded in August 1982.

99 Unspecified number of individuals (99+).

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Table 11 (cont.)

Stratigraphy Qpk Qnn Qrd Qps/Qqz Qpv

Vegetation Site2 LLI LL2 LL3 LL4 LL5

Vertebrate Quadrat IEOI IE02 IE03 IE04 IE05

Variegated Fairy-wren 2

White-winged Fairy-wren 2024 13 3425 10Brown SonglarkRufous SonglarkAustralian SittellaWhite-browed Tree-creeperMistletoebird 2 5 2

Striated Pardalote 2Brown Honeyeater - I

Black HoneyeaterPied Honeyeater 2 4

White-plumed HoneyeaterSinging Honeyeater 10 4 6

Yellow-fronted Honeyeater 6 3 3White-fronted Honeyeater - - 81 - -2.2 - - 40 - 231 2 15

Yellow-throated Miner - - 7 - 15 83927 2 - 22

Spiny-cheeked Honeyeater 210 - 4 I 3 17 12

Crimson Chat - 20 43 11

Zebra Finch - 22 -

Magpie-larkBlack-faced Woodswallow 338 2 - - I 6 - 17 1Little WoodswallowMasked WoodswallowWhite-browed WoodswallowPied Butcherbird 2 3 - 1 311 3

Grey ButcherbirdAustralian MagpieGrey CurrawongSpotted BowerbirdLittle Crow -lQ -Corvus spp - I 1: - 2

TOTAL SPECIES/SEASON 5 9 12 3 4 8 7 4 13 10 10 10 262421

TOTAL SPECIES/QUADRAT 17 12 22 19 40

Each quadrat was sampled for five days over three seasons.See Appendix l.Opportunistic sightings (not on quadrats).Total observations during F79 (February 1979), M80 (May 1980) and A81 (August 1981).

T Tracksx No season assigned for opportunistic sightings.2 All sightings of birds in flight.

* Recorded in August 1982.99 Unspecified number of individuals (99+).

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Table 12 Bird species known from the Laverton-Leonora and Sandstone-Sir Samuel Study Areas that we didnot encounter on our quadrats at Erlistoun (E) and Wanjarri (W). Sources include our ownopportunistic records from Table II (this publication) (0), the RAOU Atlas of Australian Birds(Blakers et al. 1984) (a), Anon. (1993) (b), Moriarty (1972) (m), Pianka & Pianka (1970) (p) andStorr (1985, 1986) (s). (*) denotes unconfirmed records.

NON-PASSERINES Red-necked Avocet mB1ack-throated Grebe m Bush Stone-Curlew 0

Hoary-headed Grebe 0 Gull-billed Tern bAustralian Pelican a Whiskered TernLittle Black Cormorant s Purple-crowned LorikeetLittle Pied Cormorant 0 Regent Parrot mWhite-faced Heron 0 Princess Parrot mPacific Heron 0 * Elegant Parrot mGreat Egret a Scarlet-chested Parrot mStraw-necked Ibis Major MitchelliEs Cockatoo sBlack Swan m Black-eared Cuckoo mMountain Duck 0 BoobookOwl mGrey Teal 0 Barn Owl mBlack Duck 0 Tawny Frogmouth 0

Pink-eared Duck 0 Spotted Nightjar mWood Duck 0 Rainbow Bee-eater mMusk Duck mBlack-shouldered Kite mBlack Kite m PASSERINESSquare-tailed Kite m Welcome Swallow aBlack-breasted Kite m Tree Martin 0

Whistling Kite a Fairy Martin mBrown Goshawk m White-backed Swallow 0

Little Eagle 0 Western Wedgebill aSpotted Harrier 0 Grey Fantail a* Marsh Harrier Grey-crowned Babbler mPeregrine Falcon m Banded Whiteface mAustralian Hobby 0 Western Flyeater aMalleefowl m Redthroat aLittle Button-Quail m Splendid Fairy-Wren mSpotless Crake m Rufous-crowned Emu-Wren mBlack-tailed Native-hen 0 Striated Grasswren 0

Dusky Moorhen Spinifex-bird mEurasian Coot 0 Rufous Tree-creeper pAustralian Bustard 0 Red-browed Pardalote mBlack-fronted Plover 0 Purple-gaped Honeyeater mOriental Plover s Black Honeyeater 0

Red-kneed Plover s White-plumed Honeyeater 0

Red-capped Plover 0 Grey Honeyeater mAustralian Dotterel m Orange Chat mCommon Sandpiper 0 White-fronted Chat aMarsh Sandpiper s Magpie-lark 0

Sharp-tailed Sandpiper s White-browed Woodswallow 0

Wood Sandpiper b Little Woodswallow 0

Black-winged Stilt 0 Australian Magpie 0

Banded Stilt a Australian Crow m

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The more mobile birds (non-passerines) were poorly represented in our samples; overall,51 (j{ of known species were recorded, but only 24% were recorded on quadrats (Table 3). Thequadrat-based strategy of survey was expected to be more effective for sampling relativelysedentary species with smaller home ranges and therefore greater densities in suitable habitat.Overall, 75170 of the passerine species previously known from the Study Area were recordedduring our field work, and 61!Jo of these were recorded on quadrats (Table 3). In comparison,sampling was restricted to just 10 quadrats representing only eight of the 70 surface-typesrecognised from the geological maps (sce Tables I and 2).

Each quadrat was sampled daily over five days during each of the three survey periods. Thebirds recorded on the quadrats arc listed in Table 11. Combined, the species richness of theten quadrats did not change much from season to season; a total of 44 (33 passerine) specieswere recorded in summer 1979, 46 (30) in autumn 1980, and 47 (35) in winter 1981. Insummer, 670 (539 passerines) individual birds were recorded on the quadrats; autumn andwinter sampling yielded 896 (714) and 1061 (868) bird sightings respectively. Overheadflocks of Budgerigars, Galahs, White-fronted Honeyeaters and Masked Woodswallowsaccounted for 33% of these observations, and reflected the unusually high rainfall during theperiod of our survey. Seasonal differences could be accounted for by the number of White­fronted Honeyeaters seen in winter (268).

Quadrats were classified according to the composition of their passerine assemblages (Figure15). The data were sufficient to show a consistent relationship between the quadrats' surfacestratigraphy and their passerine composition - a separation of saline, sandy, and alluvialsurfaces that is similar to the relationships noted for the other vertebrate groups. The greatermobility of the birds probably explains the different assignment of IW04, the ecotonalquadrat. There was a permanent pool of water adjacent to IW04 that attracted a variety ofbirds from adjacent Qpc habitats to the east and north.

Despite the strong patterns in species composition, we did not find significant correlationsbetween vertebrate richness and either floristic richness or number of vegetation strata. Suchcorrelations were found in the Kurnalpi-Kalgoorlie Study Area (McKenzie et al. 1992), in theNullarbor region (McKenzie and Robinson 1987) and in the Wheatbelt (Kitchener 1982,Kitchener et al. 1980). The Sandstone-Sir Samuel and Laverton-Leonora Study Areas arelocated in a region where ephemeral plants form a greater proportion of the plant assemblages,and where vegetations tend to be lower in stature and have fewer distinct strata, than theircounterparts in the more seasonal, less arid areas to the south. Pianka (1989) has summarisedsome of the factors that bear on reptile richness in such regions.

REFERENCES

Anon. (1993). Observations. W.A. Bird Notes 65,2-3.

Austin, M.P. (1984). Problems of vegetation analysis for nature conservation. In: Survey Methods for NatureConservation. (eds: K. Myers, CR. Margules and I. Musto) Vo!. I, pp 101-130. CSIRO Division of Waterand Land Resources, Canberra.

Beard, l.S. (1980). Aeolianlandforms, Western Australia. (Map at scale of 1:3,000,000 as part of VegetationSurvey of Western Australia). Vegmap Publications, Perth.

Belbin, L. (1989). PATN user's guide. CSIRO Division of Wildlife and Ecology, Canberra.

Belbin, L. (1991). Semi·strong Hybrid Scaling: a new ordination algorithm. 1. Veg. Sci. 2,491-496.

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Biological Surveys Committee of Western Australia (1984). The Biological Survey of the Eastern Goldfields. PartI: Introduction and Methods. Rec. West. Aust. Mus. Supplement No. 18,1-19.

Blakers, M., Davies, SJJ.F. and Reilly, P.N. (l984).The Atlas of Australian Birds. Royal AustralasianOrnithologists Union. Melbourne University Press, Carlton.

Buchler, E.R. (1976). A chemiluminescent tag for tracking bats and other small nocturnal animals. 1. Mammal57, 173-176.

Burbidge, A.A. and McKenzie, N.L. (1989). Patterns in the modern decline of Western Australia's vertebratefauna: causes and conservation considerations. Biological Conservation 50, 143-198.

Corben, C (1991). ANABAT ll. Bat cal/ analysis system, Version 1.1. Titley Electronics, NSW.

Curry, PJ. (1979). Long-toed Stints, Sanderling and other waders at Lake Violet, Central WA in midsummer1977-78. West. Aust. Nat. 14,109-113.

Czekanowski, J. (1932). Coefficient of racial likeness, und durschschnittliche differenz. Anthrop. ANZ. 9, 227­249.

Dell, J. and How, R.A (1988). Vertebrate Fauna. In: The Biological Survey of the Eastern Goldfields of WesternAustralia. Part 5: Edjudina-Menzies Study Area. Rec. West. Aust. Mus. Supplement No. 31, 38-68.

Dell, 1. and How, R.A (1992). Vertebrate Fauna. In: The Biological Survey of the Eastern Goldfields of WesternAustralia. Part 6: Youanmi-Leonora Study Area. Rec. West. Aust. Mus. Supplement No. 40, 20-32.

Fullard, J., Koehler, K., Surlykke, A. and McKenzie, N.L (1991). Echolocation and flight morphology ofinsectivorous bats in southwestern Australia. Aust. 1. Zoo!' 39,427-438.

Gambold, N. and Woinarski, J.C.z. (in press). Distribution patterns of herpetofauna in monsoon rainforests ofthe Northern Territory, Australia. Aust. 1. Eco!.

Henry-Hall, NJ. (1990). Nature conservation reserves in the Eastern Goldfields, Western Australia (southerntwo-thirds of CTRC System 11). Unpublished report submitted to the EPA Red Book task force. EnvironmentalProtection Authority: Perth, Western Australia.

How, R.A., Dell, J. and Humphries, W.F. (1987). The ground vertebrate fauna of coastal areas between Busseltonand Albany, Western Australia. Rec. West. Aust. Mus. 13,553-574.

How, R.A. and Dell, J. (1992). Vertebrate Fauna. In: The Biological Survey of the Eastern Goldfields of WesternAustralia. Part 7: Duketon-Sir Samuel Study Area. Rec. West. Aust. Mus. Supplement No. 40, 90-103.

Kitchener, DJ. (1982). Predictors of vertebrate species richness in nature reserves in the Western Australianwheatbelt. Aust. Wildl. Res. 9, 1-7.

Kitchener, DJ., Chapman, A., Dell, J., Muir, B.G. and Palmer, M. (1980). Lizard assemblage and reserve sizeand structure in the Western Australian wheatbelt - some implications for conservation. BiologicalConservation 17, 25-62.

McKenzie, N.L. (1984). Biological surveys for nature conservation by the Western Australian Department ofFisheries and Wildlife: A current view. In: Survey Methods for Nature Conservation. (eds: K. Myers, CR.Margules and L Musto) VoL 2, pp 88-117. CSIRO Division of Water and Land Resources, Canberra.

McKenzie, N.L and Robinson, A.C. (eds) (1987). A Biological Survey of the Nul/arbor Region, Australia.South Australian Department of Environment and Planning, Adelaide.

McKenzie, N.L, Rolfe, J.K. and Carter, D. (1987). Reptiles and Amphibians. In: A Biological Survey of theNul/arbor Region, Australia. (eds: N.L McKenzie and AC. Robinson) pp 179-210. South AustralianDepartment of Environment and Planning, Adelaide.

McKenzie, N.L and Start, AN. (1989). Structure of bat guilds in mangroves: environmental disturbance anddeterminism. In: Patterns in the Structure of Mammalian Communities. (eds: D.W. Morris, Z. Abramski, BJ.Fox and M.R. Willig), pp 167-178. Spec. PubL Mus. Texas Tech. Univ. No. 28. Lubbock, Texas.

McKenzie, N.L, Robinson, A.C and Belbin, D.L (l99Ia). Biogeographic survey of the Nullarbor district,Australia. In: Nature Conservation: Cost Effective Biological Surveys and Data Analysis. (eds: Margules,CR. and Austin, M.P). pp 109-126. CSIRO Division of Wildlife and Ecology, Canberra.

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McKenzie, NL, Belbln, L, Kelghery, G.J. and Kenneally, K.F (1991 b). Kimberley ramforest communities:patterns of species composition and Ilolocene biogeography. In: Kimberley Rainforests. (eds: N.L McKenzie,R.B. Johnston and PG. Kendriek) pp 423··51 Surrey Beatty and Sons, New South Wales.

McKenzie, N.L, Rolfe, J,K, and Youngson, W,K. (1992), Vertebrate fauna, In: The Biological Survey of theEastern Goldfields of Western Australia, Part 8, The Kurnalpi Kalgoorlie Study Area, Rec. West. Aust, Mu,\',Supplement No, 41,37-64,

Moriarty, T.K. (1972) Birds of Wanjarri (27"35', 120°40') Emu, 72, 1-7.

Pianka, E,R. (1969). Habitat specifieity, speeiation and species density in Australian desert lizards, Ecol. 50,498-502.

Pianka, E,R, (1989). Desert Lizard diversity: additional comments and some data. Am, Nat, 134, 33~··364,

Pianka, H,D. and Pianka, ER, (1970) Bird censuses from desert localities in Western Australia, Emu. 70, 17··22.

Storr, G.M. (1985). Birds of the Mid-eastern Interior of Western Australia, Rec. West. Ausl. Mus, SupplementNo. 22.

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ACKNOWLEDGEMENTSWe are grateful to the Curator of the Western Australian Herbarium for making available

facilities generally, and to KF. Kenneally and members of the technical staff for aiding theprocessing of the specimens,

We record our appreciation to the Director and staff of the W,A, Museum for theirassistance with identification and curation of the vertebrate specimens, In particular, A,Baynes, N, Cooper, J, Henry, DJ, Kitchener, LA. Smith and G,M, Storr.

Assistance with the identification of the plants was provided by TE.H. Aplin, M,I.H,Brooker, R. Carolin, RJ. Chinnock, BJ. Conn, D.A. Cooke, HJ. Eichler, J.W, Green, HJ.Hewson, S.D, Hopper, KF. Kenneally, N.S. Lander, DJ. McGillivray, TO, Mcfarlane, N,G.Marchant, B.R. Maslin, K.R. Newbey, G. Perry, B.L Rye, P,S, Short, H.R. Toelken, M,Trudgen, J.G. West, P. Weston and P.G. Wilson.

R.A. How, J. Dell and KR. Newbey provided useful discussion, The Australian Bureau ofMeteorology, Perth, provided the climatic data. We thank Tom Smith, Ken Cashin, DougWatkins and Alex Harris for assistance with the prolonged and often uncomfortable fieldprogram and, in the case of Ken Cashin, for help with data compilation,

85